Variation in nectar–sugar profile of Anchusa and allied
genera (Boraginaceae)
MASSIMO NEPI
1
*, FEDERICO SELVI
2
and ETTORE PACINI
1
1
BIOCONNET, Biodiversity and Conservation Network, Department of Environmental Sciences ‘G.
Sarfatti’, University of Siena, Via Mattioli 4, 53100 Siena, Italy
2
Department of Plant Biology, University of Florence, Via La Pira 4, 50121, Florence, Italy
Received 17 September 2009; accepted for publication 10 February 2010
The nectar–sugar profile (fructose, glucose and sucrose) of 14 species of Anchusa and five members of the allied
genera Anchusella, Cynoglottis, Hormuzakia and Lycopsis (Boraginaceae: tribe Boragineae) was determined. Most
of the species examined (c. 74%) produce sucrose-dominant nectar, whereas the remaining taxa produce sucrose-
rich nectars. Little variation in nectar–sugar composition was found in some species, even when sampling was
repeated in different years and/or localities. Average sucrose concentration was 57.75% (coefficient of variation
19.1%). The only floral morphological character that was correlated with the nectar–sugar profile is the length of
the corolla tube, as taxa with relatively long floral tubes produce nectar with lower glucose concentrations. The
flowering period is also related to sugar composition, as nectar of late-flowering species contains lower sucrose
concentrations. However, small differences in sugar profiles do not reflect phylogenetic relationships based on
molecular studies. It would appear that dry habitats and time of flowering are the main determinants of
nectar–sugar composition in the genus Anchusa sensu lato. © 2010 The Linnean Society of London, Botanical
Journal of the Linnean Society, 2010, 162, 616–627.
ADDITIONAL KEYWORDS: Boragineae – flower morphology – nectar–sugar composition – pollinators.
INTRODUCTION
Nectar is a complex mixture of substances dissolved
in water and is a nutritive source for a large
number of animal species (Nicolson, 2007; Nicolson
& Thornburg, 2007), although in some instances its
function may be much more than a simple, floral
reward (Thornburg, 2007; Nepi & Stpiczyn ´ ska,
2008). Sugars (mainly glucose, fructose and sucrose)
are the major solutes found in floral nectar. In an
extensive study of the nectar–sugar composition of
765 species of angiosperms, Baker & Baker (1983a)
found that sucrose was present in 89% of the
species examined and 84% had nectar containing all
three sugars. It is generally accepted that there
exists a co-evolutionary relationship between the
proportion of sugars in nectar and the dietary
preferences of pollinators and that this, in turn, has
led to convergence in sugar–nectar composition in
unrelated species visited by the same type of polli-
nator (Baker & Baker, 1983a, b; Nicolson & Thorn-
burg, 2007). Flowers pollinated by long-tongued
bees, butterflies, moths, hummingbirds and Old
World bats tend to produce sucrose-rich nectars,
whereas those pollinated by flies, short tongued-
bees, perching birds and New World bats tend to
produce hexose-rich nectars (Baker & Baker, 1983b).
However, it has been demonstrated that deducing
pollination syndromes from nectar types is often too
simplistic (Barnes, Nicolson & van Wyk, 1995). Phy-
logenetic conservatism governing nectar–sugar com-
position has also been demonstrated for several
angiosperm taxa, such as Lamiaceae, Ranuncu-
laceae, Brassicaceae (Baker & Baker, 1983a; Petani-
dou, 2005), Alooideae (van Wyk et al., 1993),
Papilionoideae (van Wyk, 1993) and Proteaceae
(Nicolson & van Wyk, 1998). Within these broad *Corresponding author. E-mail: massimo.nepi@unisi.it
Botanical Journal of the Linnean Society, 2010, 162, 616–627. With 3 figures
© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 616–627 616
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